外源脱落酸对醉马草内生真菌共生体幼苗建植过程的影响

doi:10.11686/cyxb2019570

摘要/Abstract

摘要： 醉马草为多年生禾本科草本植物,广泛分布于我国西北干旱-半干旱草原,其与内生真菌形成的互惠共生体具有较强的抗逆性,在自然生境中具备一定的竞争力。种子萌发及幼苗生长是植物生活史最重要的时期,决定植物未来的竞争力。通过发芽和盆栽试验,以携带(endophyte-infected, E+)和未携带(endophyte-free, E-)内生真菌的醉马草为材料,设置5个不同脱落酸(ABA)浓度,研究外源喷施ABA对醉马草内生真菌共生体幼苗建植过程的影响。结果表明:2.0 mg·L^-1的ABA浓度对醉马草种子的发芽率、胚芽长、胚根长有显著的促进作用;1.0 mg·L^-1 的ABA浓度较CK组对醉马草的净光合速率、根冠比、根表面积、根直径、根体积和根头数有显著的促进作用,但当ABA浓度为4.0 mg·L^-1时表现为抑制作用。上述结果说明内生真菌具有对外源ABA的利用能力,但这与外源ABA的浓度有很大关系。

关键词: 醉马草, 内生真菌, 脱落酸, 种子萌发, 幼苗生长

Abstract: Achnatherum inebrians is a perennial bunchgrass of the Poaceae family, widely dispersed in the arid and semi-arid grasslands of northwest China. It forms a reciprocal symbiosis with Epichloё endophyte. The presence of the endophyte provides A. inebrians strong resistance to various stresses, making it more competitive. Both seed germination and seedling growth are important stages of plant growth, determining the future competitiveness of plants. In the present study, a gradient of five different abscisic acid (ABA) concentrations was set up to explore how exogenous ABA affects endophyte-infected (E+) and endophyte-free (E-) A. inebrians seed germination and seedling growth. It was found that the germination rate, embryo length and radicle length of both E+ and E- A. inebrians seeds were significantly promoted when treated with 2.0 mg·L^-1 ABA solution. The net photosynthetic rate, root:shoot ratio, root area, root diameter, root volume and number of roots of the seedlings were higher when treated with 1.0 mg·L^-1 ABA than those in the CK group. In contrast, the germination A. inebrians seeds and seedling growth were inhibited when treated with 4.0 mg·L^-1 ABA solution. These results indicate that the interaction between Epichloё endophyte and an appropriate concentration of ABA improves seeds germination and seedling growth, and enhances the photosynthetic capacity of A. inebrians.

Key words: drunken horse grass (Achnatherum inebrians), Epichloё endophyte, abscisic acid, seed germination, seedling growth

崔雪莲, 夏超. 外源脱落酸对醉马草内生真菌共生体幼苗建植过程的影响[J]. 草业学报, 2020, 29(7): 70-80.

CUI Xue-lian, XIA Chao. Effect of exogenous abscisic acid on seedling establishment of Epichloё gansuensis-Achnatherum inebrians symbiont[J]. Acta Prataculturae Sinica, 2020, 29(7): 70-80.

参考文献

[1] Siege M R, Latch G M, Johnson M C. Fungal endophytes of grasses. Annual Review of Phytopathology, 1987, 25(1): 293-315.
[2] Saikkonen K, Gundel P E, Helander M. Chemical ecology mediated by fungal endophytes in grasses. Journal of Chemical Ecology, 2013, 39(7): 962-968.
[3] Song H, Nan Z B, Song Q Y, et al. Advances in research on Epichloё endophytes in Chinese native grasses. Frontiers in Microbiology, 2016, 7: 1399.
[4] Li C J, Nan Z B, Paul V H, et al. A new Neotyphodium species symbiotic with drunken horse grass (Achnatherum inebrians) in China. Mycotaxon, 2004, 90(1): 141-147.
[5] Nan Z B, Li C J. Neotyphodium in native grasses in China and observations on endophyte host interaction//Paul V H, Daprich P D. Proceedings of 4th international Neotyphodium grass interactions symposium. Soest: University of Paderborn, 2000: 41-50.
[6] Liu T Y, Cao M H, Li G Z, et al. Diagnosis and treatment of animal drunken horse grass poisoning. Progress in Veterinary Medicine, 2006, 27(2): 116-117.
刘图雅, 曹敏慧, 李国忠, 等. 动物醉马草中毒的诊断和治疗. 动物医学进展, 2006, 27(2): 116-117.
[7] Xia C, Christensen M J, Zhang X, et al. Effect of Epichloё gansuensis endophyte and transgenerational effects on the water use efficiency, nutrient and biomass accumulation of Achnatherum inebrians under soil water deficit. Plant and Soil, 2018, 424(1/2): 555-571.
[8] Ren A, Gao Y B, Wang W, et al. Photosynthetic pigments and photosynthetic products of endophyte-infected and endophyte-free Lolium perenne L. under drought stress conditions. Frontiers of Biology in China, 2006, 1(2): 168-173.
[9] Xia C, Li N N, Zhang X Y, et al. Role of Epichloё endophytes in defense responses of cool-season grasses to pathogens: A review. Plant Disease, 2018, 102(11): 2061-2073.
[10] Matsukura K, Shiba T, Sasaki T, et al. Dynamics of Neotyphodium uncinatum and N-formylloline in Italian ryegrass, and their relation to insect resistance in the field. Journal of Applied Microbiology, 2014, 116(2): 400-407.
[11] Zhang X X. Research progress of improved resistance of the grass to the heave mental stress by endophyte. Pratacultural Science, 2014, 31(8): 1466-1474.
张兴旭. 内生真菌提高禾草耐重金属胁迫的研究进展. 草业科学, 2014, 31(8): 1466-1474.
[12] Alboresi A, Gestin C, Leydecker M T, et al. Nitrate, a signal relieving seed dormancy in Arabidopsis. Plant, Cell & Environment, 2005, 28(4): 500-512.
[13] Ali-Rachedi S, Bouinot D, Wagner M H, et al. Changes in endogenous abscisic acid levels during dormancy release and maintenance of mature seeds: Studies with the Cape Verde Islands ecotype, the dormant model of Arabidopsis thaliana. Planta, 2004, 219(3): 479-488.
[14] Anuradha S, Rao S R. Application of brassinosteroids to rice seeds (Oryza sativa L.) reduced the impact of salt stress on growth, prevented photosynthetic pigment loss and increased nitrate reductase activity. Plant Growth Regulation, 2003, 40: 29-32.
[15] Pattanagul W. Exogenous abscisic acid enhances sugar accumulation in rice (Oryza sativa L.) under drought stress. Asian Journal of Plant Sciences, 2011, 10: 212-219.
[16] Brady S M, McCourt P. Hormone cross-talk in seed dormancy. Journal of Plant Growth Regulation, 2003, 22(1): 25-31.
[17] Fang Y, Wu J Y, Sun W C, et al. Inducing effects of exogenous ABA on seed germination and cold tolerance of winter rape seedlings. Agricultural Research in Arid Areas, 2014, 32(6): 70-74.
方彦, 武军艳, 孙万仓, 等. 外源ABA 浸种对冬油菜种子萌发及幼苗抗寒性的诱导效应. 干旱地区农业研究, 2014, 32(6): 70-74.
[18] Xu L X, Yu J J, Han L B, et al. Photosynthetic enzyme activities and gene expression associated with drought tolerance and post-drought recovery in Kentucky bluegrass. Environmental and Experimental Botany, 2013, 89: 28-35.
[19] Peleg Z, Blumwald E. Hormone balance and abiotic stress tolerance in crop plants. Current Opinion in Plant Biology, 2011, 14(3): 290-295.
[20] Clay K. Effects of fungal endophytes on the seed and seedling biology of Lolium perenne and Festuca arundinacea. Oecologia, 1987, 73(3): 358-362.
[21] Ruan Y H, Dong S K, Liu L J. Effects of exogenous abscisic acid on physiological characteristics in soybean flowering under drought stress. Soybean Science, 2012, 31(3): 385-388, 394.
阮英慧, 董守坤, 刘丽君. 干旱胁迫下外源脱落酸对大豆花期生理特性的影响. 大豆科学, 2012, 31(3): 385-388, 394.
[22] Xie J J, Wang X, Cai J, et al. Effect of exogenous application of abscisic acid and jasmonic acid at seedling stage on post-anthesis drought stress and physiological mechanism in wheat. Journal of Triticeae Crops, 2018, 38(2): 221-229.
谢静静, 王笑, 蔡剑, 等. 苗期外源脱落酸和茉莉酸减缓小麦花后干旱胁迫的效应及生理机制. 麦类作物学报, 2018, 38(2): 221-229.
[23] Zhang Z L, Zheng C X, Kang Y H, et al. Research progress of abscisic acid regulation underlying water transport in SPAC system. Water Saving Irrigation, 2016, 9: 143-147, 150.
张志亮, 郑彩霞, 康银红, 等. 激素ABA在SPAC水分传输中调节作用的研究进展. 节水灌溉, 2016, 9: 143-147, 150.
[24] Hose E, Steudel E, Hartung W. Abscisic acid and hydraulic conductivity of maize roots: A study using cell- and root-pressure probes. Planta, 2000, 211(6): 874-882.
[25] Morse L J, Faeth S H, Day T A. Neotyphodium interactions with a wild grass are driven mainly by endophyte haplotype. Functional Ecology, 2007, 21: 813-822.
[26] Hamilton C E, Bauerle T L. A new currency for mutualism? Fungal endophytes alter antioxidant activity in hosts responding to drought. Fungal Diversity, 2012, 54: 39-49.
[27] Li C J, Nan Z B, Liu Y. Methodology of endophyte detection of drunken horse grass. Edible Fungi of China, 2008, 27(Supple 1): 16-19.
李春杰, 南志标, 刘勇. 醉马草内生真菌检测方法的研究. 中国食用菌, 2008, 27(增刊1): 16-19.
[28] Wei D, Wang Y R. Germination testing methods on Achnatherum splendens seeds. Pratacultural Science, 1998, 15(4): 29-32.
卫东, 王彦荣. 芨芨草种子发芽检验方法的研究. 草业科学, 1998, 15(4): 29-32.
[29] State Bureau of Quality and Technical Supervision. Forage seed test procedure germination test, GB/T 2930.4-2017. Beijing: China Standard Press, 2017.
国家质量技术监督局. 牧草种子检验规程发芽检验, GB/T 2930.4-2017. 北京: 中国标准出版社, 2017.
[30] Faeth S H, Maajo L H, Kari T S. A sexual Neotyphodium endophytes in a native grass reduce competitive ability. Ecology Letters, 2004, (7): 304-313.
[31] Hesse U, Schöberlein W, Wittenmayer L, et al. Effects of Neotyphodium endophytes on growth, reproduction and drought-stress tolerance of three Lolium perenne L. genotypes. Grass and Forage Science, 2003, 58(4): 407-415.
[32] Savchenko T, Kolla V A, Wang C Q. Functional convergence of oxylipin and abscisic acid pathways controls stomatal closure in response to drought. Plant Physiology, 2014, 164(3): 1151-1160.
[33] Jacobsen J V, Pearce D W, Poole A T, et al. Abscisic acid, phaseic acid and gibberellin contents associated with dormancy and germination in barley. Physiologia Plantarum, 2002, 115(3): 428-441.
[34] Zhang X, Zhao C Q, Huang J, et al. Effect of exogenous abscisic acid and salicylic acid on germination and physiological characteristics of wheat seed. Chinese Journal of Applied and Environmental Biology, 2014, 20(1): 139-143.
张笑, 赵纯钦, 黄静, 等. 外源脱落酸, 水杨酸对小麦种子萌发及生理特性的影响. 应用与环境生物学报, 2014, 20(1): 139-143.
[35] Mo Y R, Li P X, Ma Z F, et al. Effects of exogenous abscisic acid on the germination of pepper seeds from different varieties. Journal of Human Ecological Science, 2014, 1(4): 28-33.
莫云容, 李培欣, 马仲飞, 等. 外源ABA 对不同品种辣椒种子萌发的影响. 湖南生态科学学报, 2014, 1(4): 28-33.
[36] Huang X H, Hu X W, Xu Z H, et al. Effects of exogenous hormones on the dormancy and germination of Leymus chinensis. Acta Prataculturae Sinica, 2013, 22(5): 183-189.
黄晓辉, 胡小文, 徐宗海, 等. 羊草种子休眠和萌发的激素调控研究. 草业学报, 2013, 22(5): 183-189.
[37] Goggin D E, Steadman K J, Emery R J N, et al. ABA inhibits germination but not dormancy release in mature imbibed seeds of Lolium rigidum Gaud. Journal of Experimental Botany, 2009, 60(12): 3387-3396.
[38] Ding J H, Li Y G, Tong J H. Effect of ABA on germination of rice seeds. Crop Research, 2012, 26(4): 328-330.
丁君辉, 李耀国, 童建华. 脱落酸对水稻种子萌发的影响. 作物研究, 2012, 26(4): 328-330.
[39] Xiong M L, Dai X, Jian Y, et al. Advances in the study of abscisic acid-dependent and non-dependent signaling pathways. Genomics and Applied Biology, 2019, 45: 568-574.
熊孟连, 戴星, 简燕, 等. 脱落酸依赖的与非依赖的信号途径的研究进展. 基因组学与应用生物学, 2019, 45: 568-574.
[40] Pinheiro C, Chaves M M. Photosynthesis and drought: Can we make metabolic connections from available data? Journal of Experimental Botany, 2010, 62(3): 869-882.
[41] Endo A, Sawada Y, Takahashi H, et al. Drought induction of Arabidopsis 9-cis-epoxycarotenoid dioxygenase occurs in vascular parenchyma cells. Plant Physiology, 2008, 147(4): 1984-1993.
[42] De Battista J P, Bouton J H, Bacon C W, et al. Rhizome and herbage production of endophyte-removed tall fescue clones and populations. Agronomy Journal, 1990, 82(4): 651-654.
[43] Liu D T, Jing Y P, Li D L, et al. Research advances in plant lateral root development. Plant Physiology Journal, 2013, 49(11): 1127-1137.
刘大同, 荆彦平, 李栋梁, 等. 植物侧根发育的研究进展. 植物生理学报, 2013, 49(11): 1127-1137.
[44] Kavar T, Maras M, Kidric M, et al. Identification of genes involved in the response of leaves of Phaseolus vulgaris to drought stress. Molecular Breeding, 2008, 21(2): 159-172.
[45] Malinowski D, Leuchtmann A, Schmidt D, et al. Growth and water status in meadow fescue is affected by Neotyphodium and Phialophora species endophytes. Agronomy Journal, 1997, 89(4): 673-678.
[46] Farooq M, Wahid A, Kobayashi N, et al. Plant drought stress: Effects, mechanisms and management, sustainable agriculture. Dordrecht: Springer, 2009: 153-188.
[47] Li D, Nan H, Liang J, et al. Responses of nutrient capture and fine root morphology of subalpine coniferous tree Picea asperata to nutrient heterogeneity and competition. PLoS One, 2017, 12(11): https://doi.org/10.1371/journal.pone.0187496.
[48] Signora L, De Smet I, Foyer C H, et al. ABA plays a central role in mediating the regulatory effects of nitrate on root branching in Arabidopsis. The Plant Journal, 2001, 28(6): 655-662.
[49] Himanen K, Boucheron E, Vanneste S, et al. Auxin-mediated cell cycle activation during early lateral root initiation. The Plant Cell, 2002, 14(10): 2339-2351.
[50] Christmann A, Weiler E W, Steudle E, et al. A hydraulic signal in root-to-shoot signalling of water shortage. The Plant Journal, 2007, 52(1): 167-174.
[51] Li S W, Leng Y, Feng L, et al. Involvement of abscisic acid in regulating antioxidative defense systems and IAA-oxidase activity and improving adventitious rooting in mung bean [Vigna radiata (L.) Wilczek] seedlings under cadmium stress. Environment Science and Pollution Research, 2014, 21(1): 525-537.